SB 508 
.W5 M4 
Copy 1 



INITED STATES DEPARTMENT OF AGRICULTURE 
BULLETIN No. 1046 

Contribation from the Bureau of Plant Industry, WM. A. TAYLOR, Chief, 
in cooperation with the Kansas Agricultural Experiment Station 



Washington, D. C. 



May, 1922 



RUST RESISTANCE 
IN WINTER. WHEAT VARIETIES 



By 

LEO E. MELCHERS, Plant Pathologist, and JOHN H. PARKER, 

in Charge of Crop Improvement, Kansas Agricultural Experiment 

Station; Agents, OflSce of Cereal Investigations 



CONTENTS . 

' Page 

Scope of the Investigation 1 

Review of the Literature 2 

Nuraery Experiments 4 

Greenhouse Experiments 14 

Comparison of Nursery and Greenhouse Results 23 

Evidence of Specific Rust Resistance . . . / 24 

Agronomic Value of Kanred Wheat .'26 

Summary 27 

Literature Cited 30 




WASHINGTON 

GOVESNMENT PRINTING OFFICi:; 

1922 






LIBRARY OF CONG«gSS 
DOCUMENTS U»Vi«ION 






UNITED STATES DEPARTMENT OF AGRICULTURE 




s^J^'^^^u 



BULLETIN No. 1046 

Contribution from the Bureau of Plant Industry 

WM. A, TAYLOR, Cliief, in cooperation with the 

Kansas Agricultural Experiment Station 




jTL^'^'Lrt. 



Washington, D. C. 



May, 1922 



EUST RESISTANCE IN WINTER- WHEAT VARIETIES.^ 

By Leo E. Melchers, Plant Pathologist, and John H. Parker, in Charge of Crop 
Improvement, Kansas Agricultural Experiment Station; Agents, Office of Cereal 
Investigations. 



CONTENTS. 



Scope of the investigation l 

Review of the literature 2 

Nursery experiments 4 

Greenhouse experiments 14 

■Comparison of nursery and greenhouse results 23 



Evidence of specific rust resistance 24 

Agronomic value of Kanred wheat 26 

Summary 27 

Literature cited 30 



SCOPE OF THE INVESTIGATION. 

A project to determine the rust resistance of existing varieties of 
winter wheats and to breed new varieties for rust resistance was 
begun in 1911 at the Kansas Agricultural Experiment Station, in 
cooperation with the Office of Cereal Investigations of the United 
States Department of Agriculture. The first two years were devoted 
to preparatory work, when no infection of stem rust was produced. 
The writers ^ took charge of the work in 1913, and the data given 
herein are those obtained since that time. 

The investigation outlined in 1913 had two major purposes: (1) 
To study the rust resistance of about 130 varieties and strains of 
winter and spring wheats, particularly to the stem rust, Puccinia 
graminis tritici Erikss. and Henn.,^ in the field and in the green- 
house; and (2) to study the inheritance of rust resistance in wheat 
and to produce hybrids adapted to commercial use. 

1 Paper No. 183 of the Department of Botany and Plant Pathology and No. 136 of the Department of 
Agronomy, Kansas Agricultural Experiment Station. 

« The writers wish to acknowledge their indebtedness to Mr. Victor H. Florell and Mr. M. N. Levine, of 
the Office of Cereal Investigations, who assisted in the greenhouse studies and in other phases of the 
investigation. 

' Puccinia graminis tritici, as used in this bulletin, has reference to those strains of stem rust used in 
the experiments in 1915, 1916, and 1917. In 1915 a strain was used to which Kanred, P1066, and P1068 
were-only partially resistant, while in 1916 and 1917 strains were used to which these varieties were very 
resistant. The strains used in 1916 and 1917 may have been one or more^of the several strains which at 
present are known not to cause normal infection of these varieties. 
79251— 22— Bull. 1046 1 



2 BULLETIN 1046, U. S. DEPAETMENT OF AGEICULTUEE. 

Climatic conditions at Manhattan, Kans., are generally unfavor- 
able for the development and spread of the stem rust of wheat; the 
development of a suitable technique for the production of severe epi- 
demics in the rust nursery was therefore an essential part of these 
investigations. 

Special attention was given to those varieties which are most 
promising agronomically. 

If a variety of hard red winter wheat could be found which was. 
resistant to stem rust and suitable for Kansas conditions, breeding 
for rust resistance would be much simpler than if it becomes neces- 
sary to cross with varieties of the durum or emmer groups. 

Hayes, Parker, and Kurtzweil (13)* recently have found that " there^ 
is an indication of linkage of durum or emmer characters and rust 
resistance, since the production of rust-resistant durums or emmers in 
the F2 and F3 generations is comparatively easy and the production 
of resistant common wheats much more difficult." Moreover, the 
only known rust-resistant varieties of the emmer or durum group& 
are spring, forms, a fact which complicates the task of obtaining a 
rust-resistant winter wheat from such a cross. Winter hardiness,, 
high yield, and good milling quality also are essential for the success 
of any variety of wheat in Kansas, which increased the complexity 
and difficulty of the problem. To obtain accurate information as 
to the resistance of existing varieties of winter wheat was, therefore,, 
the first important step. 

REVIEW OF THE LITERATURE. 

Differences in the resistance of wheat varieties to rust were known 
to exist as early as 1841, when Henslow (16) observed that some 
wheats were injured less by rust than others. La Cour (23) and Little 
(26) made similar observations. Bolley (3) noted that hardy and 
stiff-stemmed varieties with smooth, fibrous leaves seem to resist 
rust for a longer time. Anderson (1) observed that hard, flinty 
wheats are more resistant than others, believing that this might be 
due to a larger proportion of silica in the plant. Cobb and Farrer 
(35) and Farrer (35) found that wheat varieties resist leaf rusts and 
stem rusts in different degrees. Hitchcock and Carleton (16) state 
that hard varieties of wheat suffer least from rust in Kansas and 
early varieties are likely to mature before being seriously injured. 
Henning (I4) and Eriksson and Henning (11) found that certain wheat 
varieties resisted different kinds of cereal rusts. 

It has long been known that some of the emmers ( Triticum dicoc- 
cum) and certain varieties of durum wheat (Triticum durum) show 
marked resistance to stem rust. Carleton and Chamberlain (9) and 
Carleton (8) called attention to this in connection with the com^ 

'The serial niunbers (Italic) in parentheses refer to "Literature cited" at the end of this bulletin. 



EUST RESISTANCE IN WINTER-WHEAT VARIETIES, 3". 

jnercial value of the durums. Carleton (8), in discussing the stem- 
rust epidemic on wheat in 1904, observed that no wheat varieties, 
with the exception of einkorn and some of the durums, spelts, and 
emmers, showed marked resistance under all conditions. He stated 
further, that during ordinary seasons when stem rust may be quite 
prevalent the hard-kerneled Russian winter wheats are considerably 
more resistant to rust than other varieties ordinarily grown. 

Bolley (4, 5), Biff en (2), and Nilsson-Ehle (33) were among the 
first to conduct wheat-breeding experiments with the definite object 
of obtaining rust-resistant varieties. No definite plan or method of 
study, however, was described until Johnson (20) explained the 
methods used for producing an artificial rust epidemic in Minnesota 
and furnished a working basis for the studies which have been made 
since in this country in breeding cereals for rust resistance. These 
methods are further described by Freeman and Johnson (12) . They 
state that certain varieties, such as Extra Squarehead in Sweden, 
American Club in England, and Rerrarf and Ward's Prolific in Aus- 
tralia, have been shown to be resistant to rust. They add, however, 
that some of these varieties can not be said to be universally rust 
resistant, as their behavior in different countries to different biologic 
forms of rust is variable. 

Field experiments have verified early observations that some of the 
durums and emmers are much more resistant than the common 
spring- wheat varieties. Stakman (37) found this to be true in both 
field and greenhouse experiments. Melchers and Parker (29, 31) 
recently have called attention to the resistance of three winter-wheat 
varieties to stem rust and leaf rust. Waldron and Clark (4^) have 
described a variety of common wheat named Kota and stated that 
it was resistant to the strain or strains of stem rust prevalent at 
Fargo, N. Dak., Brookings, S. Dak., and St. Paul, Minn., in 1918. 
These authors state that "this resistance is decidedly greater than 
that possessed by the common spring wheats and second only to the 
more resistant durum wheats." Clark, Martin, and Smith {10) 
speak of the rust behavior of varieties of durum and common wheat 
grown during the seasons of 1914 to 1919 at several field stations in 
the northern Great Plains. They state that none of the varieties of 
common wheat grown is really rust resistant, but early-maturing 
varieties have ripened before the rust has developed extensively and 
are sometimes rust escaping. Most varieties of durum wheats are 
more or less rust resistant, as compared with common wheats. Acme, 
Monad, and D-.5 are known to be especially rust resistant. In years 
of heavy rust infection these varieties have produced the highest 
yields. When grown under comparable conditions in these and 
other experiments, the D-5 variety shows the greatest resistance of 
all varieties to stem rust. 



4 BULLETIN 1046, U. S. DEPARTMENT OF AGEICULTURE. 

NURSERY EXPERIMENTS. 

METHODS AND MATERIALS. 

Because of the infrequent occurrence of natural epidemics of stem 
rust under Kansas conditions,^ it was necessary to study varietal 
resistance in a rust nursery (PL I, figures 1 and 2), following the gen- 
eral plan suggested by Johnson (20). This rust nursery was located 
near Manhattan, Kans., on land which is low and slopes slightly 
toward the south and west. Along the south side of the nursery is 
a hedge of common barberry bushes (PI. I, fig. 2). A large drainage 
ditch on the south side carried off the surplus water. Because of 
the likelihood of frequent heavy rains during the crop season, the 
rust nursery was sown in slightly elevated plats, separated by de- 
pressed alleys, which received the surplus water and carried it into 
the main ditch. The rust nursery has been sown in various ways. 
At first a plat 1 rod square was used for each variety, but it was 
found impossible to produce severe epidemics of stem rust on large 
areas under Kansara conditions. The plats, therefore, were reduced 
to a single rod row and in 1915 to 5-foot nursery rows spaced 10 
inches apart. The seeds are sown 3 inches apart in the row. A 
small hand plow was used for opening a furrow and a seeding board 
with notches at regular intervals served to obtain uniformity in 
spacing the seed. 

The spring varieties generally were sown during the last week in 
March or the first week in April in rows close to the winter-wheat 
rust nursery. 

The rust used in these experiments up to and including 1917 was 
obtained from the Minnesota Agricultural Experiment Station. In 
1914, 1915, and 1916 the urediniospores came from greenhouse cul- 
tures of Puccinia graminis tritici, but in the fall of 1917 cultures 
were used from rusted wheat plants obtained in the field. These 
were found later to be a new strain of stem rust (30). The stock 
cultures of rust which were used in these field experiments were 
cultured on Improved Turkey (Kansas No. 2382), a variety which 
has been found in these experiments to be very susceptible to stem 
rust. When the leaves produced uredinia which were sporulating 
abundantly they were used in one of two ways: (1) The leaves 
were clipped from the plants, placed in a few quarts of water, the 
urediniospores removed, and the liquid used as a spray on the wheat 
plants in the rust nursery, or (2) the potted wheat plants bearing 
uredinia were used as centers of infection in the nursery. 

' The only natural epidemic known to the -wTiters occurred in 1904. In 1915 and 1916 stem rust was very 
prevalent in many fields in Kansas and in some instances there was an appreciable loss. In 1919 stem rust 
was uniformly present in eastern and central Kansas, and although it was difficult to estimate the actual 
. injury caused by stem rust it was one of several factors which reduced the yield and quality of wheat. 



Bui. 1046,'U. S. Dept. of Agriculture. 



Plate I. 




Fig. I. — General View of the Wheat-Rust Nursery. 

The investigation of the comparative rust resistance of varieties of wheat at the Kansas Agri- 
cultural Experiment Station was conducted here. The arrows indicate tlie location of rotary 
sprayers. 




Fig. 2. — RowIOF Barberry Bushes Adjacent to the Wheat-Rust Nursery. 

Each arrow indicates the location of a rotary sprayer. 



Bui. 1046, U. S. Dept. of Agriculture. 



PLATE II. 




Fig. I. — REMOVABLE Canvas Covers Used in the Wheat-Rust Nursery 
TO Retain Moisture. 

A urediniospore sprayer is shown in operation. 




Fig. 2. — Pot Containing Rusted Seedlings from the Greenhouse. 

Used in establisliing infection centers in the wheat-rust nursery. 



RUST RESISTANCE IN WINTER-WHEAT VARIETIES. 5 

When sprays of urediniospore decoction were used in the study at 
Manhattan, Kas., a small knapsack pressure sprayer was employed. 
All sprays were applied in the evening; if possible during periods of 
moist, cloudy weather. The plants were first sprayed with water 
and then with the urediniospore spray. Sprays of urediniospores 
were found to be unreliable, however, because of the hot dry winds 
which frequently occur in Kansas during the late spring and early 
summer. In order partially to overcome this difficulty, removable 
canvas covers were placed over a wooden framework which was 
built over the plats. These covers were used the day following the 
urediniospore sprays and aided materially in retaining moisture. 
They were easily handled by one man, being unrolled from a long 
strip of wood and drawn over the nursery plats, to be fastened at 
the corners as shown in Plate II, figure 1. 

Several attempts have been made to inoculate plants in the rust 
nursery in the fall. Not only was it difficult to obtain satisfactory 
infection, but it was of doubtful value in view of the fact that it is 
questionable whether stem rust lives over winter in Kansas to any 
great extent {17). 

On account of the unsatisfactory results obtained with the uredin- 
iospore sprays, the infection-center method of obtaining an epidemic 
was tried. This method is somewhat similar to inoculating plants 
in the field, but it is much simpler and more certain to give satis- 
factory results. The inoculated seedlings are carried from the green- 
house to the plats, where they spread the infection. Numerous 
infection centers were located in each plat, so as to provide ample 
spore material (PI. II, fig. 2). 

Most of the rust cultures for these experiments were grown on 
seedlings in 2|^-inch flowerpots, two seedlings in each pot. Fre- 
quently 4-inch pots were used, as they held more seedlings and did 
not dry out so rapidly. It was found that if small galvanized-iron 
pans were placed between the rows of wheat in the rust nursery 
early in the spring and filled with pots of inoculated seedlings, a 
most successful center of infection could be established. (PI. II, 
fig. 2) . The pans were kept filled with water at all times. 

As soon as the seedlings'died or the rusted leaves no longer produced 
urediniospores, the pots were replaced with a new set. In this man- 
ner the wheat plats were continually exposed to rust infection. It 
recently has been found that if plants in the heading stage are inocu- 
lated in the greenhouse and used in place of seedlings for the centers 
of infection, their usefulness in the field continues longer than that 
of seedling plants; hence, they are much more satisfactory. It was 
found also that wherever the centers of infection were located the 
rust obtained a start and spread rapidly from the centers to all 
adjacent plants. 



6 



BULLETIN 1046, U. S. DEPARTMENT OF AGRICULTURE. 



In 1914 only two urediniospore sprays were applied, one on April 
19 and the other on May 23. Neither of these sprays was followed 
by any noticeable rust infection. Canvas covers were not used. In 
1915 urediniospore sprays were given on April 20, 21, 27, and 29, 
and on May 7, 10, 20, 22, and 31. Inoculated seedlings were trans- 
planted into the soil in vacant rows, which were left for that purpose, 
this being the first attempt to use the infection-center method. 
Canvas covers were used and a severe epidemic was caused, as shown 
by the data on rust infection. In 1916 urediniospore sprays were 
given on April 6, 13, 14, 20, 25, 26, and 29, and on May 1, 4, 16, 19, 
and 27. Canvas covers were used. In addition to these sprays the 
infection-center method was employed. A severe epidemic resulted. 
In 1917 sprays of urediniospores were given on May 23 and 29, and 
on June 6. A few hand inoculations in the field were made in the 
spring, but the efforts were mostly directed toward establishing in- 




A, 5 per cent. B, 10 per cent. C, 25 per cent. D, 40 per cent. E, 6u per cent. F, 1(W per cent. 

Fig. 1. — Scale for e.stimating rust, illustrating six degrees of rustiness used in estimating the percentage of 
stem-rust infection. The shaded spots represent rust, and the figures represent approximately the rust 
percentages computed on the basis of the maximum of surfaces covered by rust as shown in the 100 
per cent figure (F). Figure F in the diagram represents 37 per cent of actual rust-covered surface and 
is arbitrarily selected as 100 per cent. The other percentages are in terms of figure F. 

fection centers. It was evident from the results obtained that the 
latter method was sufficiently dependable to warrant the discontinu- 
ance of urediniospore sprays. 

The common barberry (Berheris vulgaris L. ; see PI. I, fig. 2), planted 
south of the rust nursery plats, furnished some aecial infection in 
1915, 1916, and 1917. Straw, bearing telia of stem rust, was placed 
around each shrub in the fall, so as to provide the necessary telio- 
spore material to infect the barberry leaves in the spring. 

The. final field notes were taken during the latter part of June or 
early in July, at the time the nursery was harvested. These included 
the percentage of stem rust, estimated in accordance with the scale 
shown in figure 1 and used by the Office of Cereal Investigations of 
the Bureau of Plant Industry, United States Department of Agri- 
culture. Notes on the type of head, plumpness of grain, and other 
characters also were recorded. 



RUST RESISTANCE IN WINTER-WHEAT VARIETIES. 7 

The varieties grown in the rust nursery included the commonly 
grown hard red winter wheats of the Crimean group, such as Turkey 
and Kharkof, and the varieties of soft red winter wheat grown in 
eastern Kansas and other soft red winter-wheat districts. Some of 
the varieties were obtained from the Office of Cereal Investigations 
of the Bureau of Plant Industry and others from the agricultural 
experiment stations of other States. The strains grown under a 
pedigree number, and so designated in Table 1, represent pure-line 
selections made by Prof. H. F. Roberts, formerly of the department 
of botany, Kansas Agricultural Experiment Station. These strains 
were turned over to the department of agronomy in 1910, and seed 
was obtained from that department when the study of wheat varie- 
ties for rust resistance was begun. Not all the varieties have been 
grown throughout the period of experiment, because some of them 
were found to be of little or no agronomic value. Some were shown 
to be extremely susceptible to stem rust, and others were eliminated 
because of complete winter killing. A small number of spring- 
wheat varieties were grown, to obtain comparative data on rust 
infection. 

BREEDING PLAT. 

Certain varieties of spring and winter wheats were grown in a 
breeding plat each year to serve as material for crossing. The 
winter-wheat varieties were sown in the fall at the time the varieties 
were sown in the rust nurser}^. Considerable space was left between 
the rows of winter wheat, to allow for seeding spring wheats for 
crossing. Occasionally a few of the spring varieties bloomed at the 
same time as the winter wheats, thereby simplifying the work of 
making the crosses. Generally, however, it was necessary to sow 
such spring varieties in the greenhouse about the first of February. 
These were transplanted to the breeding nursery in April and May, 
thus providing some of the spring-wheat plants, which were in flower 
at the same time as the winter varieties. 

Crosses have been made between Kanred (Kansas No. 2401), 
Kansas No. 2414, and Kansas No. 2415,® three closely related winter- 
wheat varieties which are resistant to leaf rust (31) and to certain 
strains of stem rust (£9, 30) ; also between Marquis, Haynes Blue- 
stem, and Preston, varieties of spring wheat which are susceptible 
to stem rust. The Fi, Fj, and F3 generations have been grown to 
maturity, and data on the inheritance of resistance to stem rust 
(Puccinia graminins tritici) have been obtained. These results, 
however, are not presented in this bulletin. 

• These varieties have been known as P762, Pl()68, and P1066, respectively. They have recently 
been given Cereal Investigations numbers as follows: C. 1. .51-16, C. I. .5879, and C. I. 5880, respectively. 



BULLETIN 1046, V. S. DEPARTMENT OF AGRICULTUEE. 



RESULTS OF THE NURSERY EXPERIMENTS. 

The percentage of stem rust found on the varieties grown in 1915^ 
1916, and 1917 and the 3-year average for all varieties grown in all 
three seasons are shown in Table 1. Notes on the quality of grain 
also are given for the years 1916 and 1917. 

The varieties are arranged according to type and are grouped as 
to the characters, winter or spring, awned or awnless, glabrous or 
pubescent glumes, and soft or semihard to hard kernel. All of the 
winter-wheat varieties with the exception of Binkel Club are common 
wheats {Triticum vulgare). The spring grains include varieties of 
common wheat, as well as durum wheat {T. durum), emmer (T, 
dicoccum), and einkorn (T. monococcum) . 

The "Identification numbers" include the ''Pedigree number,'^ 
as used by Prof. H. F. Roberts; the "Kansas number," which is an 
accession number assigned by the department of agronomy; and 
the C. I. number, used by the Office of Cereal Investigations of the 
Bureau of Plant Industry. 

Table 1. — Stevi-rust infection of wheat varieties after artificial inoculation in the nursery 
at Manhattan, Kans., in the years 1915, 1916, and 1917, together toith data on Icernel 
quality in 1916 and 1917. 

[Key to Symbols.— Identification numbers (columns 1 and 3): C. I. = Cereal Investigations, H= hybrid, 
K= Kansas, P=pedigree. Rust infection (columns 5 to 7): T=trace, Wk= winterkilled. Quality of 
kernels (columns 9 and 10): E=excellent, F=fair, F— =poor to fair. F+=a grade better than fair, F±= 
variable quality (some poor to fair, some fair to good), G=good, G— =a grade poorer than good, G+= 
a grade better than good, G ± = variable quality (some fair to good, some good to excellent), P= poor, Vp = 
very poor.] 

Group i.— Awned, Glumes Glabrous, Kernel SEMraARD to Hard. 



Season, class, and iden- 
tification number. 


Kansas 
No. 


C.L 

No, 


Varietal name. 


Stem-rust infection (per cent). 


Quality of 
kernels. 


1915 


1916 


1917 


3-year 
aver- 
age. 


1916 


1917 


1 


2 


3 


4 


5 


6 


7 


8 


9 


10 


WINTER VARIETIES. 

Common wheats: 
P647 


2356 
2357 
2368 
2370 
2376 
2382 

2383 
2385 
2386 
2388 
2390 
2391 
2398 
2399 




Japan 


Per 
cent. 

40 

25 

25 

50 

50 

50 

45 

25 

40 

25 

40 

35+ 

40+ 

45 

65 

65 

65+ 

40 

35 

25 

25+ 


Per 

cent. 
55 
68 
70 


Per 

cent. 
Wk 
Wk 
Wk 


Per 

cent. 





""m" 

55 



57.3 

50.6 

55 

35 

57.3 

67.6 

"'67.'6' 
61.3 
61.6 
57.6 
56.6 
52.6 
52 


r+ 

F+ 
F± 

F 
F+ 

P 

G± 




P651 


Russian 




P675 


Red Winter Java 
do 




P678 


75- Wk 




P693 


1665 
5592 




60 

67+ 

47 

52 

32 

45 

30 

42 

65 

60 

48 

54 

42 

38 

45+ 

85 

47 


88 
58 

Wk 
95 
80 
95- 
Tto40 
95 
98 
Wk 
90+ 
65 
78 
95 
90 
97 
85 


F- 


P706 


Improved Tur- 
key. 
Red Winter Java 


P 


P707 




P711 


.G± ; F- 


P712 






G I F- 


P717 


1437 


Crimean 


F+ G 


P721 


G G 


P722 






G- F- 


P732 






F F 


P733 






F- 1... 


P736 






G 1 P 


P737 








F- : F- 


P742 


2434 


1538 


Ulta 


F- P 


P744 




P F 


P745 


2433 


1539 


Torgova 


F- F 


P746 




F P 


P750 


2411 


1543 


Belogltna 


F- ' P 



RUST RESISTANCE IN WINTER-WHEAT VARIETIES. 



9 



Table 1. — Stem-rust infection of wheat varieties after artificial inoculation in the nursery 
at Manhattan, Kans., in the years 1915, 1916, and 1917, together with data on kernel 
quality in 1916 and 1917 — Continued. 

Group 1. — Aavned, Glumes Glabrous, Kernel Semihard to Hard— Continued. 



Group ;?.— AwNED, Glumes Glabrous, Kernel Soft. 



■Seasan, class, and iden- 
tification number. 


Kansas 
No. 


C.I. 

No. 


Varietal name. 


Stem-rust infection (per cent). 


Quality of 
kernels. 


1915 


1916 


1917 


3-year 
aver- 
age. 


1916 


1917 


1 


2 


3 


4 


5 


6 


7 


8 


9 


10 


WINTER varieties— 

continued. 

Common wheats— Con. 

P751 

P752 


2427 


1543 


' Beloglina 


Per 

cent. 
25 
40 
50 
35+ 
30+ 
30+ 
35 
20- 
40 to 70 
40 

45 
55- 

60 
60 
20+ 


Per 
cent. 

35 

40 

60 

40+ 

65+ 

67 

65+ 

40 

10 

45 

50+ 
55- 
60+ 

65 

48 

40 

42 

45+ 

62 

65- 

85- 

82 

82+ 

75 

77 

85+ 
5± 
5± 

50+ 

60+ 

78 

45 

65+ 

70+ 

85+ 

40 

60 

45 


Per 
cent. 
60 
Wk 
88 
95 
68 
50 
95 
70 
10 
85 

95 
97 

80+ 

70 

90 

50 

70+ 

88 

90 

90 

75 
Wk 

78 

50 
Wk 

85 

15 
5 to 25 

85 

78 
Wk 
Wk 

85 

97 

85 
Wk 
Wk 

95 

88 

70+ 
Wk 

95+ 

88 


Per 

cent. 

40 

""m" 

56.6 

54.3 

49 

65 

43.3 

25.0 

56.6 

63.3 
69 

65 

66.0 

36.6 

"'"57.'6' 
70.6 
66.6 
71.6 

""73.'3' 
51.6 


P 

F- 
F- 
F- 
F- 
F- 

P 

P 

G 
F+ 

F 

F- 
G- 

F- 
P 
P 
G 

F- 
P 
F- 

F+ 

G 

G 
F- 


F 


P753 






G— 


P754 






V 


P755 






P 


P757 1 ... 






P 


P758 

P759 


2419 


1544 


Beloglina 


F 
p 


P762 


2401 
2436 


5146 


Kanred 


E 


P771 

P772 


Power Fife X 
Jonathan. 


F 
G 


P773 






G— 


P774 


2418 




Power Fife X 
Jonathan. 


G + 


P877 


F 


P882 







G 


P889 


2409 
2404 
2417 


62i7 


Turkey 


G 


P935 


Bucanera 


F+ 


P951 

P967 


Scottish Rank 


40 

60+ 

45+ 

55 + 

35 

60+ 

30 

35- 

25 

45 

40+ 

25- 

60- 

30 

45 

40 

40+ 

30 

65 


P 
G 


P973 








F 


PIOOO 


2435 




Hickling 


P 


P1003 




P1008 


2428 




Victoria . 


F 


P1013 




F— 


P1036 


2412 
2422 
2415 
2414 
2416 


2479 
2478 
5879 
5880 
1543 


Romanella 

Fem or April.... 




P1038 


65 
18.3 
20 
53.3 
66 . 

"es's' 

69 

66.6 


F 

G+ 
G+ 
F- 
F- 
G- 
G- 
F- 

F 
F- 
G+ 
F- 
F+ 


F— 


P1066 


G 


P1068 




G 


P1078 


Beloglina 


F- 


P1080 


F 


PlUl 









P1119 




1 




P1128 


1 


F 


P1131 -- - - 




F 


P1134 

P1161 


2420 
2410 


1436 
1787 
5147 
1558 
5797 
6213 
6214 

6474 
1442 


Crimean 


P 


K34 


Nebraska No. 28. 
Turkey 




K570 




P 


K2048 




Alberta Red 




F 


K2101 




Red Winter 










F 


K2123 




Defiance Hard 
Winter. 












P2141 


2413 


30- 


60- 
60+ 


61.6 


G- 
F- 


F- 


K35 


Kharkof 


F 













WINTER VARIETIES. 

Common wheats: 

P63S 


2358 
2365 


"igsi" 


BinkelClub 


68 

90+ 

90- 

" 88 
95 

85 

90— 


Wk 
70 
Wk 
Wk 
Wk 
Wk 

Wk 







F 

G 

P 

F- 
F- 

P 

F 




P668 


Michigan Bronze! 

Dietz Longberry' - - - 


P 


K51 




C.I. 1945 




Lancaster 






K36 




2008 








C.I. 1973 




New Amber 






K47 




2980 


Longberry. 
Stoner 















79251— 22— Bull. 1046- 



10 



BULLETIN 1046, U. S. DEPARTMENT OF AGRICULTUEE. 



Table 1. — Stem-rust infection of wheat varieties after artificial inoculation in the nurser-y 
at Manhattan, Kans., in the years 1915, 1916, and 1917, together with data on kernel 
quality in 1916 and 1917 — Continued. 

Group 3. — AwNED, Glumes Pubescent, Kernel Soft. 



Season, class, and iden- 
tification number. 


Kansas 
No. 


C.I. 

No. 


Varietal name. 


Stem-rust infection (per cent). 


Quality of 
kernels. 


1915 


1916 


1917 


3-year 
aver- 
age. . 


1916 


1917 


1 


2 


3 


4 


5 


6 


7 


8 


9 

F+ 
F+ 
F+ 
P 
G- 


10 


WINTER VARIETIES. 

Common wheats: 

P724 


2392 






Per 
cent. 
40+ 
20+ 
25 


Per 
cent. 

50 

45 

50 

85+ 

65 


Per 
cent. 
68 . 
Wk 
65 
95+ 
Wk 


Per 
cent. 
52.6 


F 


P1071 








P1082 




1571 
3277 


Turkey 


p 


K49 




Virginia 


p 


(?) 




Velvet Chaff 



















Group |.— AwNLESs, Glumes Glabrous, Kernel Soft. 



WINTER VARIETIES. 

Common wheats: 

P718 


2389 






30 

- 80+ 


40 

40- 

65 

40- 

65+ 

65+ 

60+ 

65 

68 

60+ 

75 

75+ 

77 

60 
67 
80 
85 
90+ 

65 
95- 


95+ 
60 
70 
95 
85+ 
90+ 
80 
90 
75 
70 
90 
Wk 

Wk 

Wk 
Wk 
Wk 
Wk 
Wk 

Wk 
Wk 


55 
60 

"73'3' 
63.3 

'"m" 
■■--■■■ 

53.3 
63.3 




G 

F 
F- 

P 
F+ 
F- 
F- 
F- 

P 

F 
G- 

F 

Vp 

F- 

F+ 

F 
P 


G — 


P739 






F 


P740 


2441 


1535 


Berdiansk 


P 


P741 




85 
40- 


P 


P748 








G— 


P765 


2402 






F+ 


P871 






40- 


F+ 


P970 


2403 
2405 
2440 
2406 






F 


P980 


"62i8' 
6216 
1733 

1744 

1915 
1923 
1969 


North Allerton.. 

Zimmerman 

Currell 

Dawson Golden 


40+ 
30+ 
25+ 


F- 


P1064 


F 


P1092 


P 


K39 




K38 




Chafl. 
Early Genesee 
Giant. 






K42 






K40 




Fultz 






K52 




Michigan Amber 
Poole 






C. I. 1979 






K48 




1980 
2997 


Fultzo-Mediter- 

ranean. 
Kofod 






K46 






C. 1.3326 




Currell 



















Group 5.— AwKLESS, Glumes Pubescent. 



WINTER VARIETIES. 

Common wheats: 

P1073 

K44 



K50. 



j Jones X Red Fife 

1933 1 Jones Winter 

t Fife. 
I Mealy , 





95 


Wk 




P- 




90 


Wk 




P 




85 


Wk 




P 





Group 6.— AwNED, Glumes Glabrous (Triticum vulgare, T. durum, and Hybrids;. 



SPRING varieties. 



Common wheats: 
C.I. 2958 



H. 3X111.-.- 
H. 4X942.10. 
H. 6X2223... 



4783 
4788 
4789 



Preston (Minn. 
No. 188). 

lumillo X Pres- 
ton. 

KubankaX Pres- 
ton. 

Kubankax Blue- 
stem. 



35 


40 


85 


53 




25 


40 


80 
68 
60 


48 


G- 
G 











p 
p 

F- 

F- 



RUST EESISTANCE IN WINTER-WHEAT VARIETIES. 



11 



Table 1 . — Stem-rust infection of wheat varieties after artificial inoculation in the nursery 
at Manhattan, Kans., in the years 1915, 1916, and 1917, together ivith data on Jcernel 
quality in 1916 and 1917 — Continued. 

Group 7.— AwNED, Glumes Pubescent. 



Season, class, and iden- 
tification number. 


Kansas 
No. 


C.I. 

No. 


Varietal name. 


Stem-rust infection (per cent). 


Quality of 
kernels. 


1915 
5 


1916 


1917 


3-year 

aver- 
age. 


1916 


1917 


1 


2 


3 


4 


6 


7 


8 


9 


10 


SPRDfG VARIETIES. 

Common wheats: 

C. I. 2442 






Black Persian... 


Per 
cent. 

— 


Per 

cent. 

10 


Per 
cent. 
10 to 40 


Per 

cent. 
13 




G 




1" 





Group S.— AWNLES.S, Glumes Glabrous. 



SPRING VARIETIES. 

Common wheats: 

C. I. 1517 

C. 1.3641 

C. I. 2S73 



Ghirka Spring. . 

Marquis 

Glyndon Fife 
(Minn. No. 163), 



■ 
35+ 


50 


98 


61 




25 


65 


95 


61 




25 


45 


85 


51 





F- 

G- 

P 



Group 5.— Awnless, Glumes Pubescent. 



SPRING VARIETIES 



Common wheats 
C. I. 2S74 




Haynes Blue- 
stem (Minn. 
No. 169). 



35+ 


40 


90 


55 






Group 70.— AwNED, Glumes Glabrous. 



Durum wheats: 

H. 3yi22A12 






lumillo X Pres 

ton. 
Arnautka 






5- 
75 






G- 


C. I. 1493 








55 






F 


C. I. 14Q4 






do 


(«) 








C. I. 1443 . . . 






Gharnovka 


• 


40 






G- 


C. 1. 1513 






Beloturka 










G 


C. I. 1736 








25 
5- 


60 
6 25 


5- 
10 
5— 

T~ 
30 


30 
13 




G 


C. I. 2094 






Kubanka 


G 


C. I. 3320 






G- 


C. I. 3320... 






Monad Selection 
Pentad (D-5)... 










G- 


C. I. 3322 














G 


C. I. 5284 
















G 





















Group /;.— AwNED, Glumes Glaprous, White. 



Emmer and Einkorn: 
C. I. 1522 






White Spring 

Emmer. 
do 














C. I. 1524 








5 
T 


T 
5 








C. I. 4781 . . 






.do 








C. I. 1526 






Yaroslav emmer 
Khapli emmer... 
Common einkorn 








C. I. 4013 . . 






T 
8 


T 
Tto35 








C.I. 2433 

























o Heavy on base; on upper culms, trace. 
'• On necks. 



c Fairly heavy at base; on culms, trace, 
d Heavy on necks; on culms, trace. 



12 BULLETIIsr 1046, U. S. DEPARTMENT OF AGRICULTURE . 

The names are those which appear in the records of the Kansas 
Agricultural Experiment Station. Most of the varieties in the 
bearded, glabrous-glumed, hard red-kerneled group are very sunilar 
to the well-known Turkey and Kharkof varieties. 

Nearly all the winter-wheat varieties proved to be very susceptible 
to stem rust (PI. III). Three of the pedigreed strains, however, were 
found to be remarkably resistant. These were Kanred and two un- 
named varieties, P1066 and P1068. These three pure-line selections 
differ morphologically from Turkey and Kharkof in the greater length 
of the short awn or beak found at the tip of the outer or empty glume. 
The average length of the beak in these three varieties is considerably 
greater than in the case of Turkey, Crimean, and Kharkof.^ The va- 
riety P762 (Kansas No. 2401) was named Kanred (from Kansas Red) 
and distributed to farmers in 1914. The other two resistant strains, 
Pi 066 and Pi 068, are very similar to Kanred; in fact, the three 
strains seem to be morphologically identical. They appear to differ 
slightly in certain agronomic characters, such as yield, winter hardi- 
ness, and grain quality. The experimental data which are available 
indicate that each of these other two selections is equal to Kanred in 
yield and other agronomic qualities, although they have not been 
grown as long in plats at the agronomy farm and have not been com- 
pared at the branch experiment stations or in cooperative experiments 
with farmers. 

These three strains did not attract any particular attention in 1915, 
as they seemed as heavily rusted (40 to 70 per cent) as many of the 
other varieties, but in 1916 and 1917 very different results were ob- 
tained.® They were almost free from stem rust (PI. IV). The 
estimated infections of rust on these three varieties in 1916 were 10, 
5, and 5 per cent respectively, and in 1917 they were 10, 15, and 5 to 
25 per cent, respectively, compared to the maximum figures of 95 to 98 
per cent on other varieties in the same seasons. 

The only other variety of winter wheat which gave any evidence 
of resistance was Kansas No. 2390. The infection of stem rust on 
plants of this variety was estimated at 40 per cent in 1915, at 30 per 
cent in 1916, and as "Trace to 40 per cent " in 1917. This variety 
was much less heavily rusted than many other varieties in 1916 and 
1917, but it does not appear to be nearly as resistant as Kanred, 
P1066, and P1068. 



1 This distinguishing character was first called to the attention of the writers by Carleton R. Ball and 
J. Allen Clark, of the Office of Cereal Investigations. 

8 In the Ught of present knowledge of the existence of several biologic strains of stem rust, with differ- 
ent infection capabilities, the results in 1915 are easily explained as being due to the presence in the rust 
nursery of one or more strains of stem rust which were able to attack these varieties. 



Bui. 1046, U. S. Dept. of Agriculture. 



Plate III. 




Typical Infection of Stem Rust of Turkey Wheat. 

This represents the susceptible strain of this vaiiely used as a checlj in the wheat-rust 

nursery in 1916. 



5ul. 1046, U. S. Dept. of Agriculture. 



Plate IV. 




A Typical Plant of Kanred Wheat from the Rust Nursery in 1916. 

Note the very slight, rust infection. The other pure lines (P 1066 and P 1068) presented a 

similar appearance. 



Bui. 1046, U. S. Dept. of Agriculture. 



Plate V . 





Typical Rust Infection of i wo Varieties of Wheat in 1 91 6. 

A, Mealy, a susceptible variety of soft red winter wheat; B, Ohirka Spring wheat, also very 

susceptible. 



Bui. 1046, U. S. Dept. of Agriculture. PLATE VI. 





«^^ 




^ :: 
* ^ 




TO. a*^^ 






— ~^ 




v^ ^:* 


A 


~B 


c^ 


^D 


E 


F 


^ -X> 




' ^^ ^. 




"*• /»# 


.^.v' 


.. ."^ y 




*t* 


.^ '" ,t 




>^ ^ '^ 


.-- V ~* 


-^ 


^ -!*■ \ 


ler " "^ 


\ 


*^ ' ' \ 


', 


<» ' 


,.. '• ^ ^" 


» *\' 






. 


- -V 


^ 


-^ '^ "^ 


"' 


V - -^ 




'\ ^! ' 


w 


^— "IL-.^ 


• ■'" - ^^ 


\'>_-^ '-. 


_"~~ V ^ 


\ _^ 


V ^ '~— 


■»-» '«»'■ «^ 


-^-- ^<* 


^G 


H 


1 


J 


K 


L 












*.^v'^ 


*-5^ 




. ^ . ^ 






'■-'v«. 


> C- . ^ 


^^ , ^V, 


•Xirf. jj i, -* « 


M 


N 


^ 


p^ 


Q 


R 



Fig. I. — Kernels of Susceptible and Resistant Wheat Varieties in 1916. 







1*— >«. ••«■ 






l^j^%i% 










ii^' -^ ^ • ^ 


V <i^^ ^y 


A 


B 


c 


D 


E 




,1 <-^^x.^ 








/f 






\\\- - 


F 


G 


H 


1 


J 












K 


L 


M 


N 






Fig. 2.— Kernels of Susceptible and Resistant Wheat Varieties in 1917. 



Bui. 1046, U. S. Dept. of Agriculture. 



PLATE VII. 




UJ ^ 




D » 



RUST EESISTANCE IN WINTER-WHEAT VARIETIES. 13 

All varieties of soft red winter wheat were found to be susceptible 
to stem rust. Plate V, A shows a typical plant of Mealy, a very 
susceptible variety. 

Several of the varieties of spring wheat showed evidence of being 
resistant. Other varieties, such as Ghirka Spring (PI. V, B), were 
found to be very susceptible. Black Persian was the only variety 
of common spring wheat which showed marked signs of resistance. 

Among the durum wheats, Beloturka (C. I. No. 1513), lumillo 
(C. I. No. 1736), Kubanka (C. I. No. 2094), Monad (C. I. No. 3320), 
and D-5 (C. I. No. 3322) all proved rather resistant to stem rust. 
A hybrid between lumillo and Preston (No. 3 X 122A12) also was 
very resistant. This hybrid resembles the durum parent in type of 
head. 

All of the strains of emmer gave some evidence of being resistant, 
as did the one strain of einkorn. Khapli emmer (C. I. No. 4013), 
although slightly rusted, never showed the large linear uredinia 
which develop on susceptible varieties. 

Usually a rather close relationship is shown between the extent 
of stem rust recorded and the quality of grain produced by a given 
variety. The grain of certain varieties, however, sometimes is 
severely injured by a medium quantity of rust, while other varieties 
with a higher percentage of rust infection will produce heavier grain. 

Kanred, P1066, and P1068 were all low in quantity of rust in 1916 
and 1917, and all produced good heavy plump kernels. Factors 
other than rust infection, of course, influence grain quality, but it 
was very evident in 1916 and 1917 that the three rust-resistant 
varieties produced better grain than the similar but more severely 
rusted varieties grown in near-by rows. 

Typical kernels of resistant and susceptible varieties produced in the 
wheat-rust nursery in 1916 and 1917 are shown in Plate VI. Table 
2 presents data showing the percentage of rust infection on the 
varieties of which kernels are shown in Plate VI. The letters in the 
first column of the table correspond with those of figures 1 and 2 of 
the plate. 



14 



BULLETIN 1046, U. S. DEPARTMENT OF AGRICULTURE. 



Table 2. — Stem-rust infection on susceptible and resistant varieties of wheat grovni in the 
rust nursery at Manhattan, Kans., in 1916 and 1917. 







1916 






Identification. 


Name. 


Class. 




Tl. VI, 

fig. 1. 


Pedigree or 

other 
number. 


Rust in- 
fection. 


A.. 


i 
P75S _ .. 


Hard red winter 


Per cent. 
65 


B 


P759 

P762 

P771 

P772 

P773 

P1038 

P1064 


Kanred 


do 


40 


C 


do 


10 


D 


do 


45 


E 


do 


50 


F 


do 


55 


G 


do 


85 


H 


do 


60 


I 


P1066 


do 


5 


J. 


P1068 


do 


5 


K 


P1071 


do 


45 


L .. 


P1078 


do 


50 


M 


P1036... . - - - - ....... 


do - 


77 


N.. .. 


Check 

C.I. 1517.. 
C. I. 3322.. 
P721 


Turkey 


do 


45 


O 


Ghirka 




50 


P 


D-5 


Red durum 




Q 




Hard red winter 


30 


R 






Soft red winter 


85 











1917. 



A . 


P7.5'^ 




Hard red winter 


95 


B . . . 


P7.59 




do 


70 


C 

D 


P762 

P765 


Kanred 


do 

do 


Trace. 
90 


E 


PS77 




do 


70 


F 


P1038 




do 


85 


G 


P1034 




.....do 


70 


H 


P1086 





do 


5 


I . 


P10S8 




do 


5 


J 


P1078 




do 


85 


K 

L . 


Kans. 2048. 
Minn. 169.. 
C. 1.2094.. 
P721 


Alberta Red 

Haynes Bluestem 


do 

Hard red spring 

Durum 

Hard red winter 


88 
85 


M 

N 


Kubanka. 


10 
40 


o 


C. I. 3277. 




Soft red winter 


95 













GREENHOUSE EXPERIMENTS. 



To check the nursery results similar experiments under controlled 
conditions were conducted in the greenhouse. 



INOCULATION METHODS. 



Nearly all the varieties, both winter and spring, which were used 
in the field were inoculated in the seedling and heading stages in the 
greenhouse. Careful records of the inoculated plants of each variety 
have been kept and their behavior compared with the behavior of 
the same variety in the field. The number of plants which can be 
grown under greenhouse conditions is more or less limited, and for 
this reason conclusions from such experiments should always be 
drawn with caution. 



INOCULATION OF SEEDLINGS. 



In inoculating seedlings the first leaf was inoculated and all others 
were trimmed off. The uredinial material came from stock cultures 
grown on Improved Turkey (Kansas No. 2382), or some other sus- 



RUST RESISTANCE IN WINTER- WHEAT VARIETIES. 15 

sceptible variety. The inoculation method used was described by 
Melchers (28) and was a modification of the method proposed and 
used by Kellerman (22) and Carleton (7) in their studies of cereal 
and other rusts. This is a satisfactory method where the supply of 
rust is limited, as there is little chance for loss of spores. The spore 
supply was collected in a Petri dish and applied to the seedlings by 
means of a flattened or a rounded needle. The leaves to be inocu- 
lated were first thoroughly dampened by stroking them several 
times between the fingers previously moistened in water. Tap 
water was used with satisfactory results, although Melhus and Dur- 
rell (32) have shown that tap water may have a toxic effect on the 
germination of certain kinds of rust spores. 

In the studies in the greenhouse the seedlings were grown in 2i-inch 
pots, illustrated in figure 2. Fifteen pots, each containing 2 seed- 
lings, or 30 seedlings in all, constituted one series of each variety. 




Fig. 2. — Seedlings grown to determine rust resistance of wheat varieties under greenhouse conditions. 
Pots 1 and 2 contain seedlings of a susceptible variety used as a check; Nos. 3 to 6 contain seedlings of 
P1068. Each of the two seedlings in each pot is trimmed to a single leaf blade which is inoculated. 

In most cases it was not necessary to inoculate more than one series 
of plants, but if the results were at all doubtful a second series was 
inoculated. Hundreds of inoculations were made on seedlings of 
Kanred, Pi 066, and Pi 068. Two to four seedlings of Improved 
Turkey (Kansas No. 2382) were used as checks for each series. 

After inoculation, the seedlings were kept in damp chambers for 
48 hours, after which they were removed and placed on the green- 
house bench. Bell jars were first used, but these proved impracti- 
cable. An inexpensive and effective damp chamber, shown in 
Plate VII, was devised. It consisted of a galvanized-iron washtub 
with a pane of glass for a cover.® 

About one-quarter of an inch of water was placed in the bottom of 
each chamber, to keep the air saturated. The chambers were kept 
in the shade, so as to maintain a temperature of 50° to 70° F. In 
warm weather, a spray of water was directed upon the damp cham- 
bers to reduce the temperatures. Difficulty was experienced during 

9 Cylinders made of galvanized iron (without permanent top or bottom), with a diameter of 15 inches 
and a height of 12 inches, have been used more recently. A removable piece of glass is used for a top. If 
these cylinders stand on damp sand, sufficient moisture is supplied to keep the chamber saturated, and 
the potted seedlings do not stand in water. Damp chambers of tliis type have proved very satisfactory. 



16 BULLETIN 1046, U. S. DEPARTMENT OF AGEICULTUEE. 

the first year in carrying the cultures through the summer, on account 
of the prolonged periods of temperatures ranging from 100° to 110° F. 
It was found that sprays of water directed upon the chamber not 
only cooled the air in the greenhouse but in addition kept the tem- 
perature in the damp chambers within a few degrees of that of the 
water itself. By such means a difference of 11° C. between the tem- 
perature inside and that outside the chambers was obtained, as 
shown by some of the readings at different dates during July and 
August. 

Johnson (21) found that the maximum temperature at which ure- 
diniospores of Puccinia graminis would germinate in a normal manner 
was about 88° F., though the experiments of the writers have shown 
that temperatm-es of 80° to 95° F. do not prevent normal infection. 
However, temperatures of 65° to 70° F. are believed to be the most 
favorable for inoculations with stem rust. These methods have been 
satisfactory in every respect; 100 per cent of infection always was- 
obtained on plants of susceptible varieties, including checks, and 
it is believed the notes on resistance or susceptibility are as depend- 
able as can be obtained under greenhouse conditions. The methods 
described, however, might not give as satisfactory results with other 
cereal rusts. 

INOCULATION IN THE HEADING STAGE. 

One plant of each variety which was grown in the rust nursery was 
allowed to develop to the heading stage in the greenhouse. One seed 
was sown in each pot in October. It has been found that by main- 
taining the proper temperature, either spring or winter wheat can be 
matured and normal seed developed in the greenhouse. Hutcheson 
and Quantz (18) have shown that temperatures from 55° to 70° F. 
are best suited to this purpose. In the experiments of the writers^ 
where plants were grown to the heading stage, a night temperature of 
about 50° F. was maintained, but not infrequently the night tempera- 
ture during the early stages of growth fell as low as 35° or 40° F. 
The temperature was kept below 75° F. during the daytime whenever 
possible. 

The plants were inoculated as soon as the heads were well out of 
the boot and the neck or peduncle exposed, as illustrated in Plate VIII. 
Two to six culms of each variety were inoculated with stem rust. In 
the case of Kanred, P1066, and P1068 the inoculations included a much 
larger number of plants. Ij^ some cases the culms of these three 
varieties were reinoculated a number of times, so as to expose them 
to infection as often as possible and at different periods of development. 
The uppermost leaf blade and the sheath of each culm of all varieties 
grown were thoroughly inoculated, as were the necks, glumes, and 
awns. These plants were then placed for a period of 48 to 72 hours 
in a specially constructed galvanized-iron damp chamber, similar to 
the one described by Parker (34). This damp chamber is of sufficient 



Bui. 1046, U. S. Dept. of Agriculture. 



Plate VIII. 




Method of Inoculating Wheat Plants at the Time of Heading and 
Seedlings at the Inoculating Stage. 



Bui. 1046, U. S. Dept. of Agriculture. 



Plate IX; 




o "^ 

2 6 



^ B 



CO 


^ 


< 


O 


I 


r-; 


O 


a 


1- 

03 


5 
H 



Bui. 1046, U. S. Dept. of Agriculture. 



PLATE X. 




Bui. 1046, U. S. Dept. of Agriculture 



Plate XI. 



Check 




A \ 



t m iHHIWW 



Check 




Fig. I. — Rust Resistance of Two Varieties of Winter Wheat in 1918. 

Seedling leaves of P1036 (A) and Kanred (B), both inoculated on April 22 and photographed 
on May 11. Compared with the rusted leaves of the susceptible check (shown at the left of 
each group) these leaves are seen to be entirely free from flecks and uredinia. 




Fig. 2. — Rust Susceptibility of Two Varieties of Spring Wheat in 1917. 

Seedling leaves: C, Arnaulka (C. I. 1493), inoculated on March 29 and photographed on April 19; 
D, hybrid 3 X 122 A 12, showing normal uredinia in this stage, inoculated on April 4 and photo- 
graphed on April 27. Although known to be resistant to some strains of stem rust in the field, 
these two varieties show visible signs of infection when inoculated with the same strain of rust 
used with A and B (fig. 1, above). 

COMPARATIVE RUST RESISTANCE OF SOME WINTER AND 
SPRING VARIETIES OF WHEAT. 



BUST RESISTANCE IN WINTER-WHEAT VARIETIES. I7 

size to accommodate about 72 olants AKm,f Koif • r. . 

^'ZZfr' '" """ "''' ''"' '"P' - illustrated^ Patrx 
aoTh, w*^ K """'"'f " """^ te-^Perature within the chamber' 

ZCl Zr^"""' '^' ""'l'"^ "''"^ '° ''^'P distnbute the watTr 
evenly. An overflow pipe m the pan at the bottom of the chamber 
earned away the surplus water. The inoculated plan s were pkcej 
on mverted empty flowerpots, to avoid setting them in wlter 

RESULTS OF THE GKEENHOOSE EXPERIMENTS 

TaMe%"'v*' "''''''"'^d from greenhouse inoculations are showri in 
Table 3. Very heavy mfection always was obtained on the checks 
and the susceptible varieties, but in the case of Kanred, P1O66 and 
PI06S no uredmia appeared. Some of the spring-whkt varie?,es 
also showed only slight to moderate infectfon. ^ The pedunc es 
glumes and awns of all susceptible varieties were just as readilv 

tnree resistant varieties, if any portion of the head becomes infected 
pant ^f tre'th?er'^",°'r^^""°"' '"'^<'™'-' »"'" ^^ ^o^^Tn 

Sn„ ^u ? ? chamber and maintained under the same condi 
tW f„ tf *'"'l'.^-'™y« became heavily infected It reviden^ 
ptL ed tn Xte'rVf ?'"""'^ ^"'•^ P^«^-'' - -ed^^ wer 
durum (CI No 2n^l> IT"""' ^- ^- ^"^ ^O'^), Kubanka 
-^^^r^l^^^Z:^^- and durum 
below) ZZlt^Jlr" r.*"™ " "'^ *''"^ "^y ^y'^bols (explained 

resut o"t:Sion'llti:eZf aTh^V^"^^? '''"'^ ''''' 
regarded as a corollary to the field re^l,t^^ ""^T *""''' ^^ 

frequently may cause a different interpretatioTof he refute ^ts 
S2d ^ *r ^^'"^ ™'y ^ ''^ '''^'^Ss ^^ inocufated A vaJety 

thouX th J'^t [ u'^t"^ "'^^'^ inoculated at another time, even 
kiectn '"'^ •" '"" '"'^ ^™^^ ^^"^^ "ay show a formal 



18 



BULLETIN 1046, U. S. DEPARTMENT OF AGRICULTURE. 



Table 3. — Results of inoculating wheat varieties with stem rust in the greenhouse at the 
Kansas Agricultural Experiment Station at Manhattan, Kans., in the winter of 
1916-17. 

[Key to identification numbers (columns 1 and 3): C. I. = Cereal Investigations, H=hybrid, K= Kansas 

P= pedigree.] 



Group 1.— 


AWNED 


, Glumes Glabrous, 


Kernel Semihard to Hard. 




Kan- 
sas 
No. 


C.I. 

No. 


Varietal name. 


Stem-rust in- 
fection. 


Type ofin- 
fection.a 




Season, class, and 
identification 


Seed- 
hngs. 


Headed 
plants. 




number. 


d . 


fe'S 

ll 


tH CD 

a's 




m 

a 
a 

1 


■ 

03 03 




■WINTER VARIETIES. 

Common wheats:— 
Continued. 
P373 . . . 








30 
/30 
\30 
/30 
t30 

30 

30 
/30 
130 

30 

30 
30 
30 


30 
13 
29 
28 
30 
28 
22 
23 
30 
30 

30 
29 
30 


3 
6 


3 

6 


S 

s 
s 
s 
s 
s 
s 
s 
s 
s 

s 
s 
s 
s 
s 
s 
s 
s 
s 
s 
s 
s 
s 
s 
s 
s 
s 
s 
s 
s 
s 
s 
s 
s 
s 

R 

s 
s 
s 
s 
s 

s 
s 
s 
s 
s 
s 
s 
s 
s 
s 
s 


s 
s 

■ ■ s" " 

"s" 
s 
s 

"s" 

"s" 
s 
s 
s 

s . 
s 
s 
s 
s 
s 
s 
s 
s 
s 

"s"" 
s 

"s" 
s 
s 
s 

"s" 

R 

s 

"s" 

s 
s 

s 
s 
s 

"s" 
s 

"s" 
s 




P647 


2356 

2357 

2368 
2370 

2376 

2382 

2383 
2385 
2386 
2388 
2390 
2391 
2398 
2399 




Japan 


Purple color. 1j 




Russian 








P651 






Do. 


P675 


Red Winter Java 
do 


"3' 
3 
3 


"3' 
3 
3 


P678... 




P693 


1665 
5592 








P706 

P707 

P711... 


Improved Tur- 
key. 
Red Winter Java 


'"9' 

3 
6 
3 
3 
6 
3 
3 
3 
4 
3 
5 
3 
3 
6 
3 


■9" 


6 
3 
3 
6 
3 
3 
3 
4 
3 
5 
3 
3 
6 
3 


Plant died. 


P712... 








P717 


1437 


Crimean. 


30 1 27 




P721 




30 
30 
30 
30 
30 
30 
30 
30 
30 
30 
r30 
\8 
30 


29 
28 
30 
30 
30 
30 
26 
30 
30 
30 
30 

8 
30 
16 
30 
30 
24 
28 

6 
21 
30 


24 
30 
30 
30 
30 

27 
30 
22 

8 
30 

7 
30 
30 
28 
28 
30 




P722 








P732 








P733 . . . 






Purple color. 


P736 . . . 








P742 

P744 


2434 


1538 


ijlta. 




P745 

P746... 


2433 


1539 


Torgova 




P750 

P751 •.. 


2411 
2427 


1543 
1543 


Beloglina 

.....do 




P752.. 


3 
3 


3 
3 




P753 








f30 
\30 
30 
30 
30 

t28 

30 
C242 
f30 
i30 

30 

30 

30 

30 
30 

{1 

30 
/30 
\30 

30 
(30 
f30 
130 




P754.. 








"■3' 
6 
6 
3 


""3" 
6 
6 
3 




P755... 








Do. 


P757... 










P758 


2419 


1544 


Beloglina 




P759... 


3 

d6. 
3 


3 

3 




P762 


2401 
2436 


5146 


Kanred 




P771 


fPower Fife x 
\ Jonathan. 




P772... 


3 
3 
6 

3 
3 
3 


3 
3 
6 

3 
3 
3 




P773 










P774 


2418 




Power Fife X 
Jonathan. 




P877 


Purple color. 


P882 










P889 


2409 
2404 
2417 


6217 


Turkey 




P935 


Bucanera 

Scottish Rank... 


"3' 
3 


"3"; 
3 




P951 




P967 


6 
3 


6 
3 




P973 

























a The symbols indicate the type of infection, as explauied on page 22. 

b Under certain conditions some varieties produced a purple color surrounding the uredinia; where this 
was pronounced it is indicated in the table. 

c Since the completion of this work about 200 additional seedlings have been inoculated, with the same 
results. 

d Since these inoculations were made, many additional culms have been inoculated, with the same 
results. 



RUST RESISTANCE IN WINTER-WHEAT VARIETIES. 



19 



Table 3. — Results of inoculating wheat varieties with stem rust in the greenhouse at the 
Kansas Agricultural Experiment Station at Manhattan, Kans., in the winter of 
1916-17— Continued. 

Group ).— AwNED, Glumes Glabrous, Kernel Semihard to Hard— Continued. 





Kan- 
sas 
No. 


C.I. 
No. 


Varietal name. 


Stem-rust in- 
fection. 


Type of in- 
fection. 




Season, class, and 
identification 


Seed- 
lings. 


Headed 
plants. 




number. 


'Zt 

P 


t-i'rt 

<a CD 

II 
1 


as 




'& 
c 

s 

CO 


■a 

o . 

-of 




WINTER VARIETIES— 

continued. 

Common wheats- 
Continued. 

PIOOO 

P1003 


2435 




Hickling 


30 

30 

30 

30 

30 

30 
c242 
c212 

30 

30 

30 

30 

30 

30 

30 

30 

30 

30 

30 

30 

30 

30 
30 
30 


30 
28 
30 
29 
27 
30 


29 
30 
27 
30 
30 
28 
29 
30 
29 
30 
30 
30 
30 

30 
30 

28 


3 
3 
3 
3 

3 

3 

d6 

d6 

6 


3 
3 
3 
3 
3 
3 


6 


s 

S 

s 

S 

S 

s 

R 
R 

s 
s 
s 
g 
s 
s 
s 
s 
s 
s 
s 
s 
s 

s 
s 
s 


s 
s 
s 
s 
s 
s 

R 
R 

s 

"s" 

s 
s 
s 
s 
s 
s 
s 
s 
s 
s 

s 
s 
s 




PIOOS... 


2428 




Victoria 




P1013 






P10.36 


2412 
2422 
2415 
2414 
2416 


2479 
2478 
5879 
5880 
1543 


Romanella 

Fern or April . . . 




P103S 




P1066 




P106S 






P107S 


Beloglina 




P1080 


Plant died. 


Pllll 






3 
3 
3 
3 
3 
3 
3 
4 
3 
3 
3 

3 
3 
3 


3 
3 
3 
2 
3 
3 
3 
4 
3 
3 
3 

3 
3 
3 


Purple color. 


P1119 






Do. 


P1128 








P1131... . 




1 




P1134 


2420 
2410 


1436 

1787 
5147 
155S 
5797 
6213 
6214 

6474 


Crimean 

Nebraska No. 28 

Turkey 

Alberta Red 

Red Winter 

Defiance Hard 
Winter. 

kharkof 

Rieti 




P1161 




K3i 




K570 






K204S 






K2101 






K2123 






IC2413 






C.I. 1442.... 






(?) 



















Group 2 


— AwNED, Glumes Glabrous, 


Kernel Soft. 




■WINTER VARIETIES. 

Common wheats: 
P638 


2353 
2365 


"im 


BinkelClub 

Michigan Bronze 
Dietz Longberry 

Lancaster 

Mammoth Red.. 
New Amber 

Longberry. 
Stoner 


f30 
130 
30 
30 
37 
30 
30 

30 


13 
30 
30 
29 
34 
30 
30 

30 


3 


3 


S 

s 
s 
s 
s 
s 
s 

s 


s 

" s " 
s 
s 
s 
s 

s 


Purple color. 






P668 


3 
3 
3 
3 
3 

3 


3 
3 
3 
3 
3 

3 


Do. 


K51 


Do. 


C.I. 1945 






K36 




2008 


Do. 


C. 1. 1973 






K47 




2980 


Do. 











Group 3.— AwNED, Glumes Pubescent, Kernel Soft. 



winter varieties. 

Common wheats: 

P724 

P1071 

K49 

(?) 



2392 



Virginia 

Velvet Chaff. 



.30 


29 


3 


3 


S 


S 


30 


30 


3 


3 


s 


s 


27 


27 


3 


3 


s 


s 


30 


30 


3 


3 


s 


s 



(0 



c Since the completion of this work about 200 additional seedlings have been inoculated, with the same 
results. 

d SincJ these inoculations were made, manv additional culms have been inoculated, with the same 
results. 

e Prominent flecks preceding uredinia. 



20 



BULLETIN 1046, U. S. DEPARTMENT OF AGEICULTURE. 



Table 3. — Results of inoculating wheat varieties with stem rust in the greenhouse at the 
Kansas Agricultural Experiment Station at Manhattan, Kans., in the idnter of 
^9i6-i7— Continued. 

Group 4.— AwNLESs, Glumes Glabrous, Kernel Soft. 





Kan- 
sas 
No. 


C.I. 
No. 


Varietal name. 


Stem-rust in- 
fection. 


Type of in- 
fection. 




Season, class, and 
identification 


Seed- 
lings. 


Headed 
plants. 




number. 






3 
3 
3 
3 
3 
3 
3 

6 
3 




PI 
1 


M 




WINTER VARIETIES. 

Common wheats: 
P718 


2389 






30 
30 
30 
30 
30 
30 
30 
30 
;30 
\30 
30 
30 
30 

30 

30 
90 
37 
30 
30 

8 
30 


30 
30 
29 
28 
30 
30 
28 
30 
17 
30 
30 
30 
22 

30 

28 
90 
36 
30 

28 

8 
30 


3 
3 
3 
3 
3 
3 
3 
6 
3 


S 

s 
s 
s 

i 

s 
s 
s 
s 
s 
s 
s 

s 

s 
s 
s 
s 
s 

s 
s 


s 
s 
s 
s 
s 
s 
s 
s 
s 

"s"" 
s 
s 

s 

s 
s 
s 
s 
s 

? 

s 




P739 






Purple color. 


P740 


2441 


1535 


Berdiansk 


Do. 


P741 




P748 










P765 


2402 








P871 








P970 


2403 

2405 

2440 
2406 






Do. 


P980 


6218 
6216 
1733 

1744 

1915 
1923 
1969 


North AUerton . . 

Zimmerman 

Currell.. 








P1064 


3 
3 
3 

6 

3 
3 
3 
3 
3 

1 
1 


3 
3 
3 

6 

3 
3 
3 
3 
3 


1 




P1092... 


Do. 


K39 


Damson Golden 

Chaff. 
Early Genesee 

Giant. . 
Purple Straw . . . 
Fultz..., 




K38 




Do. 


K42 




Do. 


K40 




Do. 


K52 




Michigan Amber 
Poole 


Do. 


C.I. 1979 






K48 




1980 

2997 
3326 


Fultzo-Mediter- 
ranean. 


Do. 


k:46 






(?) 




Currell 













Group 5.— AwNLEss, Glumes Pubescent. 



WINTER varieties. 

Common wheats: 

P1073 , 

K44 



K50. 



2408 



1933 



Jones X Red Fife 
Jones Winter 

Fife. 
Mealy 



.30 


30 


3 


3 


S 


s 


30 


25 


2 


2 


S 


s 


30 


30 


3 


3 


s 


s 



Group 5.— Awned, Glumes Pubescent (Triticum vulgare, T. durum, and Hybrids). 



SPRING varieties. 



Common wheats: 
C. 1.2958 



H3 X HI.... 
H4 X 942.10. 
H5X 223.... 



4783 
4788 
4789 



Preston (Minn. 
188). 

lumillo X Pres- 
ton. 

Kuban ka X 
Preston. 

Kuban ka X 
Bluestem. 



30 


30 


3 


3 


S 


S 


30 


30 


3 


3 


S 


s 


60 


49 


3 


3 


s 


s 


30 


30 


3 


3 


s 


s 



Purple color. 
Do. 
Do. 



Group 7.— Awned, Glumes Pubescent. 



spring variety. 



Common wheat: 
C. 1.2442 



Black Persian.. 



30 


30 


2 


2 


R3 


R3 



(/) 



/ Uredinia numerous, very small. 



EUST EESISTANCE IN WINTER-WHEAT VARIETIES. 



21 



Table 3. — Results of inoculating wheat varieties with stem rust in the greenhouses at the 
Kansas Agricultural Experiment Station at Manhattan, Kans., in the winter of 
1916-17— Qontmned. 

Group 8. — AwNLEss, Glumes Glabrous. 





Kan- 
sas 
No. 


C.I. 

No. 


Varietal name. 


Stem-rust in- 
fection. 


Type of in- 




Season, class, and 
identification 


Seed- 
lings. 


Headed 
plants. 


fection. 


Remarks. 


number. 






■St3 
I- <s 


SB 

3 
3 
3 


3) 

c 


<c . 

CS (33 

""a 
W 




SPRING VARIETY. 

Common wheats; 
C.I. 1517... 






Ghirka Spring .. 


30 
36 
30 


27 
30 
30 


3 
3 
3 


1 

s ; s 
s s 

S S 


Purple color. 


C.I. 3641 






C.I. 2873 






Glvndon Fife 
(Minn. No. 163). 

















Group S.— A WNLEss/ Glumes Pubescent. 



SPRING VARIETIES. 



■Common wheat: 
C. 1.2874 



H a y n e s Blue- 
stem (Minn. 
169). 



30 


30 


3 


3 


S 


S 



GrouTp 10.— AwNED, Glume? Glabrous. 



Durum wheats: 
H3X122A12. 
C.I. 1493..., 
C.I. 1443.... 
C.I. 1513.... 
C.I. 2094.... 



C. I. 3320. 

C. I. 3322. 
C. I. 5284. 
C. I. 2228. 



lumilloX Preston 
Arnautka. . 
Gharnovka. 
Beloturka. . 
Kubanka . . 

Monad 

D-5 

Acme 

SaragoUa. . . 



30 
59 
30 
30 
29 
/30 
\30 
60 
30 
29 


29 
49 
30 

21 
30 
30 
60 
30 
?9 


2 


2 


R4toS 

R2t0R4 
R3t0R4 

RitoRi 
R2toR4 

Rs 

Rs 

r! 

RstoS 
RstoS 


Rs 

■rV" 

RJtoS 










6 


6 










3 


3 









U). 



Group 11.— AwNED, Glumes Glabrous (White). 



Emmer and Ein- 
korn: 
C.I. 



C I. 
C.I. 

c. r. 

c. r. 

('. I. 

CI. 



1524. 

4781. 
1526. 
4013. 
3109. 

2433. 
2973. 



White Spring 

emmer. 

do 

Yaroslav emmer. 
Kliapli emmer . . 
"Wild wheat" 

( T. hermonis). 
Common einkorn 
do.... 



30 


17 


3 


3 


R2 


R6 


30 


30 


3 


3 


RstoR4 


Ri 


30 


22 


3 


2 


Rs 


Ri 


30 


30 


10 


10 


Rs 


Rs 


16 

30 
30 


16 

30 
30 






S 

s 
s 

















(*)• 

(0. 

(m). 



s LTredinia uniformly small. 

A Yellowish areas surround uredinia. 

'■ -NIo uredinia; extremely resistant. 

/ 1 lominent yellowish areas surround uredinia. 



* Heavy infection on mature plant. 
I Yellow areas pronounced in some cases. 
"> Uredinia small; resistant. 



22 BULLETIN 1046, U. S. DEPARTMENT OF AGRICULTURE. 

All the varieties of winter wheat used can be placed in one of two 
classes: (1) Those which fail to show visible signs of uredinia and 
(2) those entirely susceptible. Those of the first group are desig- 
nated by the symbol R, and those of the second group by S. Among 
the spring varieties there were different types of infection, these 
being represented by the symbols R^ to Kg and by S. Following is a 
description of the types of rust infection designated by the symbols: 

EXPLANATION OF SYMBOLS USED. 

R=Extremely resistant. 

Ri=No uredinia produced. Occasionally very small indefinite flecks, dead areas, 
or yellowish blotches occur. Signs of hypersensitiveness may be present. 

R2=Very rarely the occurrence of minute uredinia (0.1 to 0.5 mm.); presence of 
flecks, dead areas, or blotches. 

R3=Numerous minute to small uredinia (0.2 to 0.75 mm.); presence or absence of 
yellowish "islands" surrounding uredinia, or the occurrence of numerous yellow 
blotched areas surrounding or adjoining uredinia. 

R4=Uredinia numerous, variable in size and number (0.2 to 0.4 mm.); flecked or 
blotched, yellow areas present; yellow areas adjoining uredinia. 

R5=Uredinia apparently quite normal in size and shape, but presence of yellowish 
green areas surrounding or adjoining some uredinia indicative of slight resistance. 

S=Showing ordinary susceptibility; uredinia large, normal, vigorous. 

The inoculations of Kanred, P1066. and P1068 both in the seed- 
ling and in the heading stage produced consistent results, indicating 
extreme resistance. 

The hard and soft wheat varieties did not differ strikingly in 
susceptibility, except in the case of Kanred, P1066, and P1068. 
The other hard red winter wheats apparently were as susceptible as 
the soft red winter wheats, although Leach (^4) found a "decided 
correlation between the hardness and softness of wheat varieties 
and their relative susceptibility to Puccinia graminis tritici-compacti.^' 

A greater variation occurred among the varieties of durum spring 
wheat. Beloturka (C. I. No. 1513) seemed to behave very much 
like the three resistant winter-wheat varieties, in that no uredinia 
were formed. In more recent work, however, in which the same 
strain of rust was used, Beloturka (C. I. No. 1513) occasionally has 
shown a few very small uredinia. vSome of the durum varieties 
show^ed more or less uniformly the same type of infection (R2 or R^) 
as in the case of Kubanka (C. I. No. 2094), Arnautka (C. I. No. 1493), 
D-5 (C. I. No. 3322), and Gharnovka (C. I. No. 1443). A Poulard 
wheat (C. I. No. 4384) manifested a high degree of resistance. 
Among the emmers there was more or less similarity in the type of 
infection. The type of infection fluctuated in both of these groups, 
showing that various factors may affect the infection results. Two . 
White Spring emmers (C. I. Nos. 1524 and 4781) showed the R, and 
R4 types of infection, while another White Spring emmer (C. I. No. 



RUST RESISTANCE IN WINTER- WHEAT VARIETIES. 23 

1526) and Khapli emmer (C. I. No. 4013) seemed to show more con- 
sistently the type designated as R3. Khapli emmer (C. I. No. 4013) 
is the only variety of spring wheat so far studied that under all 
conditions seemed to exhibit a high degree of resistance in the field 
and in the greenhouse. Even this variety, however, may be classed 
at times as R, instead of R3. The seedlings of the so-called wild 
wheat of Palestine, Triticum hermonis Cook (C. I. No. 3109), really 
a form of emmer, all proved to be susceptible. 

The purple color mentioned under "Remarks" occurred in some 
varieties and surrounded the uredinia. It does not appear con- 
sistently in any one variety. Environmental conditions seem to 
affect its production. 

COMPARISON OF NURSERY AND GREENHOUSE RESULTS. 

In general, the results of the greenhouse inoculations were similar 
to those produced under field conditions in the rust nursery. All of 
the winter-wheat varieties were found to be susceptible except 
Kanred, PIO66, and PIO68. Kansas No. 2390, which appeared to be 
partially resistant in the rust nursery, gave no evidence of being 
resistant in the greenhouse. 

In the nursery, results on rust behavior usually are obtained on 
varieties in the heading stage, but in the greenhouse the seedling 
stage is most commonly used. Determining the resistance of varie- 
ties as seedlings is the most convenient method, particularly when a 
number of distinct biologic strains of rust are being used. It perhaps 
is the most severe test that can be given a variety and should always 
serve as a check on nursery results. The results thus obtained, 
however, scarcely can be considered as the sole criteria of the actual 
field resistance or susceptibility of a variety; in fact, plants showing 
certain effects when inoculated in the seedling stage in the green- , 
house may respond very differently to the same rust when they are 
subjected to it in the heading stage under field conditions. No 
definite statements as to the cause of these differences can be made 
at this time. The same factors causing resistance or susceptibility 
may operate in all stages of growth, but the reaction of the host and 
parasite at various stages of development may be different. 

It is possible that time of maturity may have an important 
influence on the extent of the rust on a given variety, but if the rust 
is epidemic before the plants reach the heading stage there can be 
no doubt as to the plants having been exposed to infection. The 
differences perhaps are due to complex factors in the developmental 
stages in a variety, which may cause a different response to rust 
infection. To whatever cause these differences in behavior in the 
seedling and heading stages may be due, the behavior of any variety 



24 BULLETIN 1046, U. S. DEPARTMENT OF AGRICULTURE. 

under field conditions is of first importance from the agronomic and 
plant-breeding viewpoint. The growing of wheat varieties in the 
rust nursery places them under conditions at least as severe as those 
to which a commercial field is subjected in a natural epidemic. 
Field tests must finally determine the value of any variety. 

EVIDENCE OF SPECIFIC RUST RESISTANCE. 

In the studies made of the resistant varieties, Kanred, P1066, and 
P1068, it was noted that their reaction to rust infection was entirely 
different from anything that had been observed in any other variety. 
Such varieties as Khapli emmer (C. I. No. 4013), White Spring 
emmers (C.I.Nos. 1524 and 1526 and Minnesota No. 1165), and the re- 
sistant durums Kubanka (C. I. No. 2094) and Arnautka (C. I. No. 
1493) are known to show resistance in the seedling stage in the 
greenhouse as well as under field conditions. Their resistance in 
the seedling stage is shown by the formation of relatively sm;ill 
uredinia, surrounded by yellow or yellowish white areas, the occur- 
rence of minute brown or yellowish dead areas, the presence of 
yellowish islands, or other characters generally regarded as indicativ3 
of resistance or hypersensitiveness. This evidence of resistance is 
illustrated in Plate XI. All the spring- wheat varieties which the 
writers have studied and which are classed as highly resistant show 
such reactions to rust infection, and almost always very distinct 
uredinia, though frequently small, are formed in inoculated seedlings 
of these varieties. 

Kanred, P1066, and P1068 are unique in their behavior toward 
Puccinia graminis tritici, as hundreds of seedlings and of culms in 
the heading stage have been inoculated with this strain of rust and 
not a single uredinium ever has been observed. The entirely rust- 
free and unflecked inoculated leaves of Kanred and P1066 are illus- 
trated in Plate XI, fig. 1, A and B. These varieties may be said 
to be immune ^^ from this particular stem rust, if it be assumed that 
the controlled conditions provided in the greenhouse are as favorable 
and that exposure to infection is as severe as under natural field 
conditions, and that seedling inoculations are as severe a test as can 
be given to a variety. They are certainly more strikingly resistant 
to Puccinia graminis tritici than any other varieties of common 
wheat (Triticum vulgare) which have been studied by the writers. 

Because of this specific behavior these varieties have been used 
as differential hosts in separating certain biologic strains of stem 
rust of wheat. The inoculation studies with these varieties have 
been carried over a long period, including every month in the year, 

i» The word "immune" is here used to mean freedom from any macroscopic evidence of rust infection 
or to designate the inability of the rust fimgus to sporulate. 



RUST RESISTANCE IN WINTER- WHEAT VARIETIES. 25 

under various temperature conditions and at various stages of 
development of the plant. The work has been done by different 
persons at different agricultural experiment stations and always 
with the same results. The only visible evidence of infection in 
the seedling stage has been the occasional appearance of very indefi- 
nite, scarcely visible, whitish flecks, generally less than 0.1 millimeter 
in diameter. These indefinite flecks are not similar to the areas or 
flecks occurring on the seedlings of the resistant emmers and durums 
(PI. XI, fig. 2, C) and are very much less conspicuous. In this 
respect these three varieties of winter wheat are distinct in their 
behavior. 

The behavior of Kanred, P1066, and P1068 in the nursery and 
field is not greatly different from that in the greenhouse. Table 1 
shows that these varieties had very low percentages of stem-rust 
infection, varying in 1916 and 1917 from a trace to 10 per cent. 
In 1915 the percentages recorded were higher. In view of present 
knowledge of the existence of several distinct biologic strains of 
wheat-stem rusts {38, 39, 40, 4^), this rather heavy infection very 
probably was due to the use of one or more biologic strains of stem 
rust similar to, if not identical with, the one recently described by the 
writers (30). 

When mature culms of the three resistant varieties were inoculated 
in the greenhouse with cultures of Puccinia graminis tritici, a response 
on the part of the host to the rust infection was only occasionally 
visible. Slightly yellowish or brownish white minute dead areas 
were sometimes vaguely visible, indicating that infection had 
occurred but that the organism had ceased to develop. 

The results reported in this bulletin establish the fact that Kanred 
and two other very similar pure lines of hard red winter wheat are 
resistant to certain biologic strains of black stem rust. 

More recent studies {25, 30, 41) have shown that these varieties 
are not resistant to all the known strains of stem rust. Extensive 
field observations made in 1919 and 1920 have indicated, however, 
that Kanred is much less severely injured by most of the stem-rust 
strams occurring in Kansas than are Turkey and Kharkof, the other 
varieties commonly grown. Reports from Wisconsin, Alabama, 
Nebraska, New York, Illinois, Missouri, Iowa, California, and New 
South Wales (Australia) indicate that these three varieties have 
shown resistance to stem rust, while the Minnesota and the South 
Dakota agricultural experiment stations report them rather severely 
rusted, although in South Dakota Kanred showed less rust than 
Turkey. Because of the existence of distinct strains of stem rust 
it is probable that the behavior of these varieties will vary in different 
seasons and in various sections of the country. 



26 BULLETIN 1046, V. S. DEPARTMENT OF AGRICULTURE. 

RESISTANCE TO LEAF RUST. 

Observations (31) made at Manhattan, Kans., during the 5-year 
period from 1915 to 1919, inclusive, and field notes recorded in all 
sections of the State in 1919, 1920, and 1921 show that these three 
pure lines of Crimean wheat are remarkably resistant also to leaf 
rust {Puccinia triticina) as it occurs in Kansas. Mains and Jackson 
(27) also have found these three varieties to be very resistant to leaf 
rust under field conditions and where the plants were approaching 
the heading stage. According to these workers, however, seedlings 
of these varieties do not prove to be resistant to leaf rust when 
inoculated and maintained under greenhouse conditions. 

The resistance to leaf rust has been manifested also in experimental 
field sowings made in the States of Alabama, California, Missouri, 
North Carolina, North Dakota, Oregon, South Dakota, Tennessee, 
Texas, Virginia, and Wisconsin, and also in New South Wales, 
Australia. Present knowledge of the leaf-rust problem and the 
records from a wide range of sowings subjected to severe epidemics 
of leaf rust indicate rather definitely that Kanred, P1066, and P1068 
will maintain this high degree of resistance under a wide range of 
conditions. It should not be supposed, however, that the resistance 
of these varieties to leaf rust will be absolute under all conditions or 
in the presence of all the biologic strains of leaf rust which may exist. 

AGRONOMIC VALUE OF KANRED WHEAT. 

Kanred wheat presents a unique combination of desirable agro- 
nomic characters, a fact which is of even greater significance than its 
resistance to rusts. Jardine (19) described the origin and history 
of Kanred wheat and called attention to its higher yield, earliness, 
and cold resistance. 

Call and Salmon (6) state that ''at Manhattan, the average pro- 
duction of Kanred has been 4.5 bushels per acre more than Turkey 
and 4.7 bushels more than Kharkof." It has outyielded these 
varieties in every season but one and in that season (1914) practically 
equaled the others. Salmon (36) presents further experimental data 
on the superiority of Kanred and gives statements regarding the 
value of Kanred from a large number of farmers who have grown 
the new variety. 

It is estimated that at least 1,500,000 acres were sown to Kanred 
wheat in Kansas in the fall of 1920, and it is expected that within a 
few years this variety will occupy a large percentage of the hard 
winter-wheat acreage of Kansas. If the area sown to Kanred should 
reach 7, 000, 000 acres and the yield should be increased 3 bushels 
per acre, with wheat selling at $1 per bushel, other factors re- 
maining unchanged, the annual value added to the Kansas wheat 



EUST RESISTANCE IX WIXTER-WHEAT VARIETIES. 27 

crop as a result of the production of Kanred wheat would be $21,000,- 
OOO. A statement of the agronomic value of this variety will be 
found in Circular 194 of the United States Department of Agriculture. 
Although the problem of breeding wheat for resistance to stem rust 
has been greatly complicated by recent discoveries of a number of 
distinct biologic strains of rust which are present in the several grain- 
:growing regions, Kanred wheat in the future probably will prove of 
great value as a parental variety in crosses, for it certainly contains 
factors for resistance to some strains of leaf rust and stem rust. 
There is good evidence that these factors are transmitted in wheat 
hybrids in the same general way as the factors for other characters. 
Kanred wheat is being used by the Tennessee Agricultural Experi- 
ment Station as the rust-resistant parent in a series of crosses with 
adapted varieties of soft red winter wheat, in the hope of producing 
varieties of soft red winter wheat resistant to leaf rust and otherwise 
equal to the best varieties now being grown, which are often severely 
damaged by leaf rust. Kanred also has been used at the Kansas 
and Minnesota Agricultural Experiment Stations as a parent in a 
large number of crosses. It is too early to make any predictions as 
to the value of any of these hybrids, although several of them appear 

promismg. 

SUMMARY. 

(1) Field experiments to determine the resistance to black stem rust 
(Pvccinia graminis tritici) of about 100 varieties and strains of winter 
wheat, many of them pure-line selections, and of a few varieties of 
spring wheat, were conducted in a rust nursery at Manhattan, Kans., 
in 1915, 1916, and 1917. Greenhouse experiments were conducted 
during the winter of 1916-17, using the same varieties. Special 
methods were developed for producing rust epidemics under the pre- 
vailing climatic conditions of Kansas. 

(2) In the rust nursery severe epidemics were produced each 
season, and the percentage of rust infection probably represented 
the maximum rust attack which the varieties would encounter under 
field conditions. 

(3) All the winter-wheat varieties grown were found to be suscep- 
tible to stem rust except Kanred and two very similar pure-line 
selections, P1066 and P1068. These three varieties were found to be 
resistant. Another pure-line strain, Kansas No. 2390, gave evidence 
of being partially resistant. 

(4) Plumpness of kernels usually is reduced by severe rust attack. 
The three resistant varieties produced grain of good quality in 1916 
and 1917, when other varieties grown under the same conditions 
but much more severely rusted produced very badly shrunken 
kernels. 



28 BULLETIN 1046, U. S. DEPARTMENT OF AGRICULTUEE. 

(5) Several varieties of spring wheat proved rust resistant under 
the conditions of these experiments, though the Black Persian was 
the only spring-wheat variety of the common or bread-wheat group 
(Triticum vulgare) which was found to be resistant. Of the varieties 
of durum or macaroni wheat (Triticum dwum), Beloturka (C. I. No. 
1513), lumillo (C. I. No. 1736), Kubanka (C. I. No. 2094), Monad 
(D-1), and Pentad (D-5) showed definite signs of resistance to stem 
rust. A hybrid of lumillo X Preston, resembling the durum parent, 
also was found to be rust resistant. All of the strains of emmer and 
einkorn grown gave some evidence of resistance. 

(6) In the greenhouse experiments the plants were studied for 
rust resistance at two stages of growth, viz, as seedlings and at the 
time of heading. The results were very similar to those in the field 
experiments. All the winter-wheat varieties were susceptible except 
three— Kanred, P1066, and P1068. Kansas No. 2390, which ap- 
peared to be somewhat resistant in the field, showed no evidence of 
resistance at either stage of growth in the greenhouse. Most of the 
spring-wheat varieties which the field experiments had shown to be 
resistant also gave more or less evidence of resistance under green- 
house conditions. This was not true, however, of einkorn. 

(7) Although the results obtamed in the field and those in the 
greenhouse agree fairly well, final conclusions as to rust resistance of a 
variety should not be drawn from greenhouse tests alone. The com- 
bined evidence from nursery experiments and inoculations of seed- 
lings and of plants in the heading stage under greenhouse conditions 
is much more likely to agree with actual field trials, which must 
always be the final test of the practical value of any variety. 

(8) The behavior of the rust parasite on the inoculated plants of 
the three resistant varieties^Kanred, P1066, and P1068 — seems to be 
different from that in other varieties described as resistant. In most 
other varieties prominent flecks are nearly always present in 8 to 12 
days after inoculation, and most frequently small uredinia are pro- 
duced. In these three varieties, however, flecks are very rarely 
visible, and in no instance have even the most minute uredinia been 
observed. 

(9) Reports from Alabama. California, Illinois, Iowa, Missouri, 
Nebraska, New York, Wisconsin, and New South Wales indicate 
that these three varieties are resistant to stem rust; but Minnesota 
and South Dakota report them rather severely attacked by stem rust. 
The occurrence of distinct strains of stem rust complicates the prob- 
lem of predicting what their behavior may be during different seasons. 
Present knowledge of the distribution of stem-rust strains and whether 
they occur each season in definite regions is so limited that the resist- 
ance or susceptibility of those wheats in any region may differ from 
season to season. 



RUST RESISTANCE IN WINTER- WHEAT VARIETIES. 29 

(10) The very light infection of leaf rust in sowings made in Ala- 
bama, Arkansas, California, Missouri, North Carolina, North Dakota, 
Oregon, South Dakota, Tennessee, Texas, Virginia, and Wisconsin, 
and in New South Wales has proved that these three varieties are 
extremely resistant to leaf rust also. Present knowledge of the 
leaf-rust problem indicates rather definitely that these varieties will 
maintain this high degree of resistance under a wdde range of condi- 
tions. 

(11) Kanred, one of the three rust-resistant pure lines, has an" 
unusual combination of desirable characters. In Kansas it yields 
from 3 to 5 bushels more per acre than either Turkey or Kharkof, 
the varieties commonly grown. It ripens a little earlier, thus 
escaping some of the damage from drought and hot winds during the 
ripening period. Kanred also seems to be more winter hardy in 
Kansas than other varieties and survives the severe winters with 
less loss from winterkilling. In milling and baking quality it 
apparently is equal to Turkey and Kharkof, varieties of hard red 
winter wheat which have established a world-wide reputation for 
quality. 

(12) Experiments and the experience of large numbers of farmers 
have shown that Kanred is adapted to other sections of the hard 
winter-wheat area, and it is now rapidly being introduced and widely 
grown in Oklahoma, Texas, Nebraska, eastern Colorado, and some 
other States. 

(13) Kanred wheat is believed to have considerable potential 
value also as a parental variety to be used by plant breeders in com- 
bining its rust resistance and other valuable characters with those of 
the varieties of other classes of wheat adapted to the several wheat- 
growing districts. 



LITERATURE CITED. 

(1) Anderson, H. C. L. 

1890. Rust in wheat. Experiments and their objects. In Agr. Gaz. N. S. 
Wales, V. 1, pt. 1, p. 81-90, ilkis. 

(2) BiFFEN, R. H. 

1907. Studies in the inheritance of disease resistance. In Jour. Agr. Sci.^ 
V. 2, pt. 2, p. 109-128. 

(3) BOLLEY, H. L. 

1889. Wheat rust. Ind. Agr. Exp. Sta. Bui. 26, 19 p., 9 fig. 

(4) 1905. Experiments and studies upon wheat. In No. Dak. Agr. Exp. Sta. 

15th Ann. Rpt. 1904, p. 34-51, 1-5 fig. (on pi.). 

(5) 1909. Some results and observations noted in breeding cereals in a specially 

prepared disease garden. In Amer. Breeders' Assoc. Rpt.^ v. 5 
(1908), p. 177-182. 

(6) Call, L. E., and Salmon, S. C. 

1918. Growing wheat in Kansas. Kans. Agr. Exp. Sta. Bui. 219, 51 p., 11 
fig. 

(7) Carleton, Mark Alfred. 

1903. Culture methods with Uredinea?. In Jour. Appl. Micros, and Lab. 

Methods, v. 6, no. 1, p. 2109-2114. 

(8) 1905. Lessons from the grain-rust epidemic of 1904. U. S. Dept. Agr., 

Farmers' Bui. 219, 24 p., 6 fig. 

(9) and Chamberlain, Joseph S. 

1904. The commercial status of durum wheat. U. S. Dept. Agr., Bur. Plant 

Indus. Bui. 70, 70 p., 1 fig., 5 pi. 

(10) Clark. J. Allen, Martin, John H., and Smith, Ralph W. 

1920. Varietal experiments with spring wheat on the northern Great Plains. 
U. S. Dept. Agr. Bui. 878, 48 p., 2 tig., 3 pi. Publications on 
cereals in the Great Plains area, p. 48. 

(11) Eriksson, Jakob, and Henning, Ernst. 

1896. Die Getreideroste. Jhre Geschichte und Natur sowie Massregeln gegen 
dieselben. vii, 463 p., 5 fig., 13 col. pi., 1 col. tab. Stockholm. 
Litteraturverzeichnis, p. 446-457. 

(12) Freeman, E. M., and Johnson, Edward C. 

1911. The rusts of grains in the United States. U. S. Dept. Agr., Bur. 
Plant Indus. Bui. 216, 87 p., 2 fig., 1 pi. Bibliography, p. 79-82. 

(13) Hayes, H. K., Parker, John H., and Kurtzweil, Carl. 

1920. Genetics of rust resistance in crosses of varieties of Triticum vulgare 
with varieties of T. durum and T. dicoccum. In Jour. Agr. Re- 
search, V. 19, no. 11, p. 523-542, pi. 97-102. Literature cited, p. 
541-542. 

(14) Henning, Ernst. 

1894. Nagra ord om olika predisposition for rost a sad. In K. Landtbr. 
Akad. Handl. och Tidskr., arg. 33, p. 205-217. 

(15) Henslow, J. S. 

1841. Report on the diseases of wheat. In Jour. Roy. Agr. Soc. England, 
V. 2, p. 1-25. 
30 



RUST RESISTANCE IX WINTER-WHEAT VARIETIES. 31 

(16) Hitchcock, A. S., and Carleton, M. A. 

1893. Preliminary report on rusts of grain. Kans. Agr. Exp. Sta. Bui. 38, 
14 p., 3 pi. 

(17) 1894. Second report on rusts of grain. Kans. Agr. Exp. Sta. Bui. 46, 9 p. 

(18) HuTCHEsoN, T. B., and Quantz, K. E. 

1917. The effect of greenhouse temperatures on the growth of small grains. 
In Jour. Amer. Soc. Agron., v. 9, no. 1, p. 17-21, 1 fig., 2 pi. 

(19) Jardine, W. M. 

1917. A new wheat for Kansas. In Jour. Amer. Soc. Agron., v. 9, no. 6, 

p. 257-266. 

(20) Johnson, Edward C. 

1911. Methods in breeding cereals for rust resistance. In Proc. Amer. Soc 
Agron., V. 2 (1910), p. 76-80. 

(21) 1912. Cardinal temperatures for the germination of uredospores of cereal 

rusts. (Abstract.) In Phytopathology, v. 2, no. 1. p. 47. 

(22) Kellerman, W. A. 

1903. Uredineous infection experiments in 1902. In Jour. MycoL, v. 9, 
no. 65, p. 6-13. 

(23) La Cour, J. G. 

1863. Sygdommene i kornet og midlerne derimod. Zn Tidsskr. Land0kon., 
Raekke 3, Bd. 11, p. 249-264. 

(24) Leach, Julian G. 

1919. The parasitism of Puccinia graminis tritici Erikss. and Henn. and 
Puccinia graminis tritici-compacti Stak. and Piem. In Phyto- 
pathology, V. 9, no. 2, p. 59-88, pi. 4-6. 

(25) Levine, M. N., and Stakman, E. C. 

1918. A third biologic form of Puccinia graminis on wheat. In Jour. Agr. 

Research, v. 13, no. 12, p. 651-654. 

(26) Little, W. C. 

1883. Report on wheat-mildew. Jour. Roy. Agr. Soc. England, ser. 2, v. 
19, p. 634-691. 

(27) Mains, E. B., and Jackson, H. S. 

1921. Two strains of Puccinia triticina on wheat in the United States. (Ab- 
stract.) In Phytopathology, v. 11, no. 1, p. 40. 

(28) Melchers, Leo E. 

1915. A way of obtaining an abundance of large uredinia from artificial 
culture. In Phytopathology, v. 5, no. 4, p. 236-237. 

(29) and Parker, John H. 

1918. Three varieties of hard red winter wheat resistant to stem rust. (Ab- 

stract.) In Phytopathology, v. 8, no. 2, p. 79. 

(30) 1918. Another strain of Puccinia graminis. Kans. Agr. Exp. Sta. Circ. 68, 4 p. 

(31) 1920. Three winter wheat varieties resistant to leaf rust in Kansas. In 

Phytopathology. v.lO, no. 3, p. 164-171, 3 fig. Literature cited, p. 171. 

(32) Melhus, I. E., and Durrell, L. W. 

1919. Studies on the crown rust of oats. Iowa Agr. Exp. Sta. Research Bui • 

49, p. 115-144, 6 fig. BibUography, p. 143-144. 

(33) Nilsson-Ehle, H. 

1911. Kreuzungsuntersuchungen an Hafer und Weizen. II. In Lunds 
Univ. Arsskr., n. f., afd. 2, bd. 7, nr. 6, 84 p. Literaturverzeichnis 
zu der Einleitung, p. 20. 



32 BULLETIN 1046, U. S. DEPARTMENT OF AGEICULTUEE. 

(34) Parker, John H. 

1918. Greenhouse experiments on the rust resistance of oat varieties. U. S. 

Dept. Agr. Bui. 629, 16 p., 2 fig., 3 pi. Literature cited, p. 16. 

(35) Rust in Wheat Conference. 

1891-92. Report of the proceedings, 2d-3d session, 1891-92. Sydney (N. S. 
Wales), Adelaide (S. Australia). 

(36) Salmon, S. C. 

1919. Establishing Kanred wheat in Kansas. Kans. A'gr. Exp. Sta. Giro. 

74, 16 p., 7 fig. 

(37) Stakman, E. G. 

1914. A study in cereal rusts. Physiological races. Minn. Agr. Exp. Sta. 
Bui. 138, 56 p., 9 pi. BibUography, p. 50-54. 

(38) and Hoerner, G. R. 

1918. The occurrence of Puccinia graminis tritici-compacti in the southern 

United States. In Phytopathology, v. 8, no. 4, p. 141-149, 2 fig. 
(39) Levine, M. N., and Leach, J. G. 

1919. New biologic forms of Puccinia graminis. [Preliminary paper.] In 

Jour. Agr. Research, v. 16, no. 3, p. 103-105. 

(40) and Piemeisel, F. J. 

1917. Biologic forms of Puccinia graminis on cereals and grasses. In Jour 
Agr. Research, v. 10, no. 9, p. 429-496, pi. 53-59. 

(41) United States Department op Agriculture. 

1920. Kanred wheat epoch-making for Kansas ... Jn U. S. Dept. Agr. 

Weekly News Letter, v. 8, no. 6, p. 6. 

(42) Waldron, L. R., and Glare, J. A. 

3919. Kota, a rust-resisting variety of common spring wheat. In Jour. 
Amer. Soc. Agron., v. 11, no. 5, p. 187-195, pi. 7. 



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